Three cryptic species in Asecodes (Förster) (Hymenoptera, Eulophidae) parasitizing larvae of Galerucella spp. (Coleoptera, Chrysomelidae), including a new species

ore

JHR 30: 5 |—64 (20 I 3) JOURNAL OF Apeenreviewed open-access journal
ee iene (>) Hymenoptera

www.pensoft.net/journals/jhr The imernaonl Society of Hymenopeariss, RESEARCH

Three cryptic species in Asecodes (Forster)
(Hymenoptera, Eulophidae) parasitizing larvae
of Galerucella spp. (Coleoptera, Chrysomelidae),

including a new species

Christer Hansson'*, Peter A. Hambiack??

| Zoological Museum, Department of Biology, Sdlvegatan 37, SE-22362 Lund, Sweden 2 Department of
Botany, Stockholm University, SE-106 91 Stockholm, Sweden

T urn:lsid:zoobank. org:author:EC9LEABD-71 15-4B05-BC80-9195C86FA55D
* urn:lsid:zoobank. org:author: CLA8ABIC-38 1 4-45 9F-AE78-3981 CCFA417C

Corresponding author: Christer Hansson (Christer. Hansson@biol.lu.se)

Academic editor: S. Schmidt | Received 10 November 2012 | Accepted 14 December 2012 | Published 30 January 2013
urn:lsid:zoobank. org:pub:B693C4F7-D9C9-44C5-IODA-47EB253B9IBI6

Citation: Hansson C, Hamback PA (2013) Three cryptic species in Asecodes (Forster) (Hymenoptera, Eulophidae)
parasitizing larvae of Galerucella spp. (Coleoptera, Chrysomelidae), including a new species. Journal of Hymenoptera

Research 30: 51-64. doi: 10.3897/JHR.30.4279

Abstract

Three morphologically very similar species of Asecodes Forster (Hymenoptera: Eulophidae) are reviewed.
Asecodes parviclava (Thomson) is removed from synonymy under A. /ucens stat. rev., and differentiated
from A. lucens (Nees) and A. lineophagum sp. n. All three species develop as gregarious endoparasitoids in
larvae of Galerucella spp. (Coleoptera: Chrysomelidae), but each species has its own unique host range.
Asecodes lineophagum attacks only Galerucella lineola (Fabr.) and A. lucens only G. sagittariae (Gyllenhal),
whereas A. parviclava parasitizes G. tenella (L.), G. calmariensis (L.) and G. pusilla (Duftschmid). The Ase-
codes species are similar but display small though distinct morphological differences, and are distinguished
also through molecular differences. The genetic distance in mitochondrial CO1 ranged from 2.3% to
7.3% between the species. Five names, one valid and four synonyms, were available for this group of spe-
cies, but none of them was linked to a primary type. To promote stability of nomenclature, primary types
are designated for all five names, neotypes for Eulophus lucens Nees, Entedon mento Walker and Derostenus
parviclava Thomson, and lectotypes for Entedon chthonia Walker and Entedon metagenes Walker. Entedon
mento, E. chthonia and E. metagenes remain synonymized under A. lucens.

Copyright Christer Hansson, Peter A. Hambdck. This is an open access article distributed under the terms of the Creative Commons Attribution License
3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

52. Christer Hansson & Peter A. Hambéack / Journal of Hymenoptera Research 30: 51-64 (2013)

Keywords
CO1, koinobiont endoparasitoids, host specificity, neotype designation, lectotype designation

Introduction

Species of Asecodes Forster are described in the literature as parasitoids of chrysomelid
beetles (Askew and Viggiani 1978, Kamijo 1986, Schauff 1991). Asecodes lucens
(Nees) (as A. mento (Walker)), was reared from Galerucella nymphaeae (L. ) (Hippa
and Koponen 1984) and Lochmaea suturalis (Thomson) (Golightly 1962) (Coleoptera:
Chrysomelidae). In an unpublished study in Sweden the junior author and associates
also reared A. /ucens from five additional species of Galerucella, G. calmariensis (L.),
G. lineola (Fabr.), G. pusilla (Duftschmid), G. sagittariae (Gyllenhal) and G. tenella
(L.). However, parasitism rates differed considerably in different Galerucella species
collected from the same locality. These patterns caused us to wonder whether a single
species, A. /ucens, was using multiple hosts or if there were populational differences in
host range. To examine the possibility that population differentiation occurs within A.
lucens based on host use we sequenced both mitochondrial and nuclear genes (Ham-
back et al. unpubl data). This analysis indicated the occurrence of multiple species.

In this paper, we identify morphological traits that separate three species of Ase-
codes, corresponding also to molecular and biological data. We compared the morpho-
types with previous descriptions and found that two correspond to described species,
A. lucens (Nees) and A. parviclava (Thomson), whereas the third was not described.
The new species is described under the name A. lineophagum.

Material and methods

The colour photos were made with a Nikon SMZ 1000 stereomicroscope and a Nikon
DS-5M camera. To eliminate reflections from the metallic and shiny body, a dome-
light manufactured as described by Kerr et al. (2008), was used as the light source for
photography. Photos were taken at different focus levels and Helicon Focus Pro version
4.75 was used to merge them into a single image. The photography of wing interfer-
ence patterns is described in detail by Shevtsova and Hansson (2011). The SEM pho-
tos were made from uncoated specimens on their original cardboard mounts. ‘These
were taken in low vacuum mode on a JEOL JSM 5600LV SEM microscope.

Morphological terms follow Gibson (1997). For illustrations of the morphological
terms see www.neotropicaleulophidae.com.

The genetic distances were estimated from pairwise comparisons of 784 base pairs
corresponding to the three prime end of CO1 (as used in Hambiack et al., unpub-
lished). In the analysis, 7 individuals of A. /ucens, 23 individuals of A. parviclava and 3
individuals of A. lineophagum were used. The analysis was done in PAUP* ver. 4.0.a125
(Swofford 2002), and genetic distances were calculated under the Kimura-2-parameter

Three cryptic species in Asecodes (Forster) (Hymenoptera, Eulophidae) parasitizing larvae... 53

model (K2P) with pairwise deletion of missing data. Additional mitochondrial and
nuclear genes data were used for reconstructing the phylogeny (Hambick et al., un-
published), but were not included in the estimate of genetic distances due to a less
extensive data set than for CO1.

Museum acronyms

BMNH. the Natural History Museum, London, United Kingdom (N. Dale-Skey
Papilloud)

CH private collection of Christer Hansson

GNM the Natural History Museum, Gothenburg, Sweden (C.G. Jonsson)

LUZM Lund University Zoology Museum, Sweden (R. Danielsson)

NHRS the Natural History Museum, Stockholm, Sweden (H. Vardal)

ZSM Zoologische Staatssammlung, Miinchen, Germany (S. Schmidt)

Designation of primary types

Five names were available for this complex of species prior to this investigation, with
one name, A. /ucens, considered as valid and four other names as synonyms under the
latter name. However, none of the names were fixed to a primary type and three of the
names lacked type material altogether. For nomenclatural stability names must be fixed
to a type specimen, and we therefore designate primary types, neotypes or lectotypes,
for all five available names.

Neotypes

Eulophus lucens was described by Nees from a single female caught on Robinia pseudoa-
cacia in Sickershausen [Bavaria, Germany] June 22, 1812, which was placed in his own
collection. Specimens in the collection of Nees no longer exist apart from specimens
sent to Westwood, now in the Oxford University Museum of Natural History (Gra-
ham 1988). No material of E. lucens exists in the Oxford collection (Graham 1988)
and it is thus safe to assume that the holotype of E. lucens was destroyed. A female
from Hungary, Vas County, Szakonyfalu, collected 23.vi.1960, agrees well with the
description of E. /ucens and is here designated as neotype. The locality in Hungary is
the closest to the type locality where it has been possible to find material of this species.

Entedon mento was described by Walker from an unspecified number of males
from near London and from Belfast. There is no material that agrees with the original
description, either in the general collection or in the type collection of the Natural His-
tory Museum, London, where the material Walker based his descriptions is kept. In
the type collection, the box supposed to contain some type material of E. mento is emp-

54 Christer Hansson & Peter A. Hambéack / Journal of Hymenoptera Research 30: 51-64 (2013)

ty, but with a note “mento??”. Thus, it appears the material on which Walker based
his description of E. mento is now lost. A female from England, Middlesex, Southgate,
collected 6.vi.1972, fits the description of E. mento and is here designated as neotype
for Entedon mento. ‘The species was allegedly described from males, but males and fe-
males of this species are very similar. Further, Walker frequently misidentified the sex
(Graham 1963), and a female is selected for neotype to make this species comparable
to the other species of this complex, all of which are represented by females.

Derostenus parviclava was described by Thomson from an unspecified number of
females he collected on Oland, an island in the Baltic Sea, and by G.F. Moller in Hol-
meja, a locality in Skane, the southernmost province in Sweden. The collection of C.G.
Thomson is in the Lund museum, and the collection of G.F. Moller is in the Natural
History Museum in Gothenburg, both in Sweden. However, neither collection has any
specimens from the type localities of D. parviclava. There is a female under the name
D. parviclava in the G.F. Moller collection from Bokeberg (labeled “Bok”), which is
a locality very close to Holmeja, one of the type localities. Because Thomson was very
specific concerning localities from the province Skane, where he lived and worked, it
seems unlikely that he interchanged “Holmeja” with “Bokeberg”. However, the female
from Bokeberg agrees well with the original description of D. parviclava and it is from
a locality very close to one of the original type localities. This specimen fulfills the cri-
teria for a neotype for Derostenus parviclava and is designated as such here.

Lectotypes

The descriptions of Entedon chthonia and E. metagenes do not have information on the
number of specimens used, and neither has been fixed to a primary type. The type col-
lection of the Natural History Museum, London, has a specimen each of E. chthonia
(type no. 5.2603) and E. metagenes (type no. 5.2604). These specimens fit the original

descriptions and are here designated as lectotypes.

Biology of the parasitoids

The Asecodes species included here are gregarious koinobiont endoparasitoids of beetle
larvae (Stenberg and Hamback 2010). Females lay their eggs in the early larval stage
and successful parasitoid development leads to a mummification of the host larva. The
host larval mummies are black and morphologically resemble larvae, whereas unpara-
sitized larvae form soft yellow pupae typical of chrysomelid beetles (photos in Hamback
2004). After successful development the parasitoid larvae pupate inside the mummified
host larva. The number of parasitoid pupae in the mummies is highly variable, from
1-14 within one mummy. The emergence of adult parasitoids typically occurs in inter-
vals, which probably reflects separate egg-laying events and thus indicates superparasit-
ism as a common trait. The number of parasitoids in each host affects both the sex ratio

Three cryptic species in Asecodes (Forster) (Hymenoptera, Eulophidae) parasitizing larvae... 55

and the adult body size of emerging offspring. Single emerging parasitoids are invariably
females, but at high parasitoid densities the sex ratio is male biased (up to 80%) (Sten-
berg and Hambick 2010). It also seems that density dependence in sex ratio and adult
body size are correlated with host species, as indicated by a comparison of parasitoids
emerging from Galerucella tenella and G. calmariensis (Stenberg and Hamback 2010).
Parasitism rates may at times be very high, close to 100%, but may at other times
be very low. In Sweden, where this study was performed, there seems to be a latitudinal
shift in parasitism rates, at least for some hosts. ‘The parasitism rates for G. calmariensis
and G. tenella in northern localities, close to Umea, are typically very high, between
50% and 100%, but less than 10% in more southern localities. The genus contains
both strictly monophagous species and oligophagous species and the different Galeru-
cella species often occur in the same localities, but on different wetland plants. There
are often large differences in the parasitism rates between Galerucella species within the
same locality. For instance, parasitism rates may be very high on G. /ineola and very
low on other Galerucella species in one locality, whereas parasitism rates are high in
another species in another locality. The different parasitism rates are not likely to be
due to phenological differences or spatial distributions within localities because host
plants colonised by different larval species may occur on neighboring plant individuals.

Genetic analysis

Genetic distances of A. lucens vs A. lineophagum calculated from mitochondrial gene
data varied between 4.8—6.0 % (mean = 5.3%), A. lucens and A. parviclava 5.3-7.3 %
(mean = 6.4%) and A. lineophagum vs A. parviclava 2.3-3.8% (mean = 3.0%). Varia-
tion was estimated as 2.6% for A. parviclava, whereas no variation was found within

A. lucens and A. lineophagum for the sampled individuals.

Identification

For identification of the species treated here the following additions can be made to the
latest key to European species of Asecodes in Askew and Viggiani (1978).
Couplet 9, replace “mento” with “11”, and include the following:

11 Forewing speculum open (Fig. 2); propodeal callus with 3—5 setae (Fig. 22).
yp ace va tee atten yen tisdt ata teen wn bt wncay pata a ous bane Rae A, lineophagum sp. n.
- Forewing speculum closed posteriorly by costal setal line (Fig. 6); propodeal

Calls wathe2-setac: Mle? 3) ei aees Rosset ween een anette ah aetna cement teh saretaee ct 12
12 Forewing bare just behind marginal vein, and relatively sparsely setose (Figs
AON SCE ASO RIC LZ to vate cvecdaowtenceunroucensqscsestecedos aves A. parviclava (Thomson)

— Forewing setose just behind marginal vein setose, and relatively densely setose

Cas eRe ote o Ba RS iret iy hte ieee etre eee rit a ere er re eee A. lucens (Nees)

56 Christer Hansson & Peter A. Hamback / Journal of Hymenoptera Research 30: 51-64 (2013)

Species treatments

Asecodes lineophagum sp. n.
urn:lsid:zoobank.org:act:3B302B7C-F323-45E3-8E78-19C080201E1A
http://species-id.net/wiki/Asecodes_lineophagum

Figures 30 eo G17, 2 2526.27,

Diagnosis. Forewing (Fig. 2) with speculum open posteriorly (i.e. setal line absent),
bare just behind marginal vein and otherwise relatively sparsely setose; propodeal callus
with 3—5 setae (Fig. 22).

Description. FEMALE. Length 1.0-1.8 mm.

Antenna dark brown (Fig. 10). Frons below frontal suture metallic purple with
upper-lateral corners close to eyes and frontal suture golden-green (Fig. 26), above su-
ture golden green. Vertex metallic purple inside ocellar triangle, golden-green outside
triangle. Mesoscutum black with metallic purple tinges (Fig. 16). Scutellum metallic
bluish-green (Fig. 16). Axillae black with metallic purple tinges (Fig. 16). Dorsellum
metallic bluish-green (Fig. 16). Propodeum metallic bluish-green (Fig. 16). Coxae,
femora and tibiae dark brown to black, and shiny (as in Fig. 1); fore tarsus dark brown,
mid and hind tarsi with tarsomeres 1-3 yellowish-white, tarsomere 4 dark brown.
Forewing hyaline (Fig. 2), wing interference pattern as in Fig. 3. Petiole dark brown
to black. Gaster with 1* tergite metallic bluish-green, remaining tergites dark brown to
black with metallic purple tinges.

Antenna as in Fig. 10. Frons below frontal suture with weak reticulation, above
suture smooth; antennal scrobes join on frontal suture. Vertex with very weak reticula-
tion inside ocellar triangle, smooth outside triangle.

Mesoscutum with weak reticulation (Fig. 17). Scutellum with very weak reticula-
tion in anterior 2/3 (Fig. 17), posterior 1/3 smooth. Axillae with weak reticulation
(Fig. 17). Dorsellum slightly convex and smooth (Fig. 17). Propodeum with a wide
groove along anterior margin (Fig. 17), with weak reticulation; propodeal callus with
3—5 setae. Forewing (Fig. 2) bare just behind marginal vein, speculum open, setation
relatively sparse, and with 5—9 admarginal setae.

Petiole as a short, transverse, narrow stripe. Gaster circular.

Ratios. Height of eye/malar space/width of mouth = 2.6/1.0/1.8; shortest distance
between posterior ocelli/posterior ocellus and eye/posterior ocellus and occipital mar-
gin = 10.4/5.4/1.0; width of head/width of mesosoma = 1.1; length of forewing/length
of marginal vein/height of forewing = 2.4/1.0/1.0; length of postmarginal vein/length
of stigmal vein= 0.6; length of mesosoma/length of gaster = 1.0.

MALE. Length 0.9-1.4 mm.

Very similar to female except antenna (Fig. 13) with scape wider, flagellomeres
longer and more slender, and apical two flagellomeres distinctly separated.

Ratios. Height of eye/malar space/width of mouth = 2.1/1.0/1.6; length of meso-
soma/length of gaster = 1.0—1.2.

Three cryptic species in Asecodes (Forster) (Hymenoptera, Eulophidae) parasitizing larvae... 57

Hosts and sex ratio. All Swedish specimens were reared from Galerucella lineola
(Coleoptera: Chrysomelidae) on Salix spp., mainly S. cinerea. The sex ratio of each
clutch on average is closer to one than for A. /ucens, and the standard deviation is
distinctly higher in A. neophagum. Ratio female/male (n = 48): 3.1042.68/2.2742.17

Material examined. HOLOTYPE female (BMNH) labelled “SWEDEN: Up-
pland, Ludden, 59°46'18"N, 18°40'19"E, 21.vi.2011, ex Galerucella lineola on Sa-
lix cinerea”. PARATYPES. 459 493(BMNH, CH, LUZM, NHRS, ZSM): 259
193 with same label data as holotype; 32. 1d“Sweden: Uppland, Flaktan, 59°46'54"N,
17°44'30"E, 24.vii.2011, ex Galerucella lineola on Salix cinerea.”; 32. 535“SWEDEN:
Uppland, Liljekonvaljholmen, 59°48'18"N, 17°39'51"E, 18.vi.2011, ex Galerucella li-
neola on Salix sp.”; 62 43“SWEDEN: Uppland, Mortsjén, 59°38°39” N 18°09°58” E,
5.vii.2011, ex Galerucella lineola on Salix cinerea’; 13“SWEDEN: Uppland, Sundan-
gen, 59°33.954'N, 16°51.292'E, 25.vi.2011, ex Galerucella lineola on Salix cinerea’;
39 6d“SWEDEN: Uppland, Haknis, 59°43'05"N, 17°41'57"E, 25.vi.2011, ex Ga-
lerucella lineola on Salix cinerea”; 19 “SWEDEN: Skane, Skaralid, 6—17.viii.1994, M
. Sporrong”; 32. (on two pins) labelled “Smaland” [which is: SWEDEN: Smaland,
without further information]; 19 14¢“SWEDEN: Skane, Sdvde, 5.vii.1985, C. Hans-
son”; 1¢“SWEDEN: Smaland, Hyltebruk, 7—14.ix.1986, J. Ardé”; 1¢“NORWAY:
Jostedalen, Gupne, 19.vii.1979, Hull University Expedition”.

Asecodes lucens (Nees)
http://species-id.net/wiki/Asecodes_lucens
Figures 6-8, 12, 15, 20, 21, 23, 24

Eulophus lucens Nees, 1834: 175. Neotype female in ZSM, designated here.

Entedon mento Walker, 1839: 28. Neotype female in BMNH, designated here. Syn-
onymized by Graham (1993: 227).

Entedon chthonia Walker, 1839: 122. Lectotype female in BMNH, designated here.
Synonymized by Graham (1993: 227).

Entedon metagenes Walker, 1848: 230. Lectotype female in BMNH, designated here.
Synonymized by Graham (1993: 227).

Asecodes lucens (Nees), Graham (1993: 227).

Diagnosis. Forewing (Figs 6, 8) with speculum closed posteriorly by a setal line, setose
just behind marginal vein and otherwise relatively densely setose; propodeal callus with
desctaetPie.23)

Hosts and sex ratio. All Swedish specimens were reared from Galerucella sagit-
tareae (Coleoptera: Chrysomelidae) on Lysimachia thyrsiflora, L. vulgaris, and Poten-
tilla palustris. The sex ratio of each clutch is female biased. Ratio female/male (n = 48):
4.27+1.83/0.92+0.94.

Material examined. TYPE MATERIAL: Neotypes of FE. lucens (ZSM)
and E. mento (BMNH), lectotypes of E. chthonia (BMNH) and £. metagenes

58 Christer Hansson & Peter A. Hambéck / Journal of Hymenoptera Research 30: 51-64 (2013)

(BMNH), all types are females. ADDITIONAL MATERIAL: DENMARK: 49
23(CH, LUZM). HUNGARY: Vas Co. 52 24(ZSM). SWEDEN: Skane 1069
13¢(swept) (BMNH, CH, LUZM); Uppland 2052 44¢(919 25¢from Lysi-
machia, 11492. 193 from Potentilla palustris) (BMNH, CH, NHRS, ZSM); Oland
14 (swept) (CH).

Asecodes parviclava (Thomson), stat. rev.
Figures 1, 4,5, 9, 11, 14, 18, 19, 25

Derostenus parviclava Thomson, 1878:272—273. Neotype female in GNM, designated here.
Asecodes parviclava (Thomson), Boucek and Askew (1968: 131).
Synonym of Asecodes lucens (Nees) (Graham 1993: 227).

Diagnosis. Forewing (Figs 4, 9) with speculum closed posteriorly by a setal line, bare
just behind marginal vein and otherwise relatively sparsely setatose; propodeal callus
with 2 setae (as in Fig. 23).

Hosts and sex ratio. The Swedish specimens were reared from Galerucella cal-
mariensis and G. pusilla (Coleoptera: Chrysomelidae) on Lythrum salicaria, and G.
tenella on Filipendula ulmaria. The number of samples is smaller than for the other two
species (n = 23, 10 from G. calmariensis, 6 from G. pusilla, 7 from G. tenella). Ratio
female/male: 2.09+1.16/0.91+0.95.

Material examined. TYPE MATERIAL: Neotype female of D. parviclava (GNM).
ADDITIONAL MATERIAL: HUNGARY: Vas Co. 12 (BMNH); SWEDEN:
Skane 219 164(CH, LUZM); Uppland 489 214(249 8¢from G. calmariensis, 109
8d from G. pusilla, 142 53from G. tenella) (BMNH, CH, NHRS); Vastergétland
142 53(CH, LUZM).

Discussion

The separation of A. /ucens into three species based on molecular and morphological
evidence is supported by biological data. Asecodes lucens and A. lineophagum were
reared from only one host species, Galerucella sagittariae and G. lineola, respectively,
whereas A. parviclava was reared from three host species, G. tenella, G. calmarien-
sis and G. pusilla. These observations have also been confirmed with independent
observations of parasitoid behaviour in the laboratory, where females were found
to attack the respective host species, but not other species (L. Fors, unpubl. data).
The delimitation of three species is also supported by observations in the field,
where one species of Galerucella larvae may be heavily parasitized and another is
not attacked in the same locality. Such parasitism patterns have, however, not been
observed for G. tenella, G. calmariensis and G. pusilla. In fact, earlier studies show

Three cryptic species in Asecodes (Forster) (Hymenoptera, Eulophidae) parasitizing larvae... 59

Figure |. Asecodes parviclava (Thomson), female habitus (length = 1.5 mm).

strong correlations in parasitism rates between G. tenella and G. calmariensis among
localities (Hambick et al. 2006). Moreover, field observations suggested that para-
sitism rates on G. tenella were consistently higher when this species was sympatric
with G. calmariensis. The reason for this pattern was not resolved (Hambiick et al.
2006), but the pattern suggest that the parasitoid population may mediate indirect
interactions between its hosts, as is known for other host-parasitoid systems. The
current information on the species delimitation within Asecodes was important to
identify pairs of host species where such effects would be likely. Based on previous
information, we could have expected similar indirect effects also for other species
pairs but the novel information on population differentiation among parasitoid in-
dividuals suggest this not to be the case.

60 Christer Hansson & Peter A. Hambick / Journal of Hymenoptera Research 30: 51-64 (2013)

speculum

without setae, +
i.e. speculum open Se.

with setae
speculum

with setae here, *.
i.e. speculum closed

Figures 2-9. Asecodes spp.,wings, females 2=3 A. lineophagum sp. n. 2 transparent wings 3 wing interfer-

ence pattern 4=5 A. parviclava (Thomson) 4 transparent wings 5 wing interference pattern 6=7 A. lucens
(Nees) 6 transparent wings 7 wing interference pattern 8 A. /ucens, forewing, neotype 9 A. parviclava,
forewing, neotype.

In view of our findings, the previous host records of Galerucella nymphaeae and Loch-
maea suturalis for A. lucens need confirmation. Investigation of Asecodes specimens reared
from these hosts might quite possibly reveal additional cryptic species in this group.

Three cryptic species in Asecodes (Forster) (Hymenoptera, Eulophidae) parasitizing larvae... 61

Figures 10-15. Asecodes spp., antennae 10-12 females 10 A. lineophagum sp. n. II A. parviclava
(Thomson) 12 A. ducens (Nees) 13-15 males 13 A. Lineophagum 14 A. parviclava 15 A. lucens.

62 Christer Hansson & Peter A. Hambick / Journal of Hymenoptera Research 30: 51-64 (2013)

meas

Figures 16-23. Asecodes spp. 16—21 thoracic dorsum, females 16=17 A. ineophagum sp. n. 18-19 A.

parviclava (Thomson) 20-21 A. lucens (Nees) 22=23 lateral propodeum in side view (anterior part to the
left), female 22. A. lineophagum 23 A. lucens.

Three cryptic species in Asecodes (Forster) (Hymenoptera, Eulophidae) parasitizing larvae... 63

Figures 24-27. 24-25 Asecodes spp., neotypes, females 24 A. lucens (Nees) 25 A. parviclava (Thomson)
26-27 A. lineophagum sp. n., head in frontal view 26 female 27 male.

Acknowledgements

Thanks to museum staff (names listed under the museum acronyms above) for loan of
type material, to the Department of Biology, Lund University, for the use of their SEM
facility, and to Elisabeth Weingartner and Johannes Bergsten for assisting in handling
the material. The study was funded by a grant from the Swedish Research Council
Vetenskapsradet (contract no. 621-2009-4943).

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Boucéek Z, Askew RR (1968) Palearctic Eulophidae, excl. Tetrastichinae. Index of Entomopha-
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Gibson GAP (1997) Morphology and terminology. In: Gibson GAP, Huber JT, Woolley JB
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